Structure and dynamics of Japanese beech (Fagus japonica Maxim.) stools and sprouts in the regeneration of the natural forests

Structure and spatial distribution of stools and root-collar sprouts of Japanese beech (Fagus japonica) were studied to clarify the regeneration processes of the stool and the population, and the ecological importance of this stool formation in five quadrats of the natural forests with different forest floor vegetation on the Pacific side of Japan. F. japonica dominates in the canopy of each quadrat. Most of sprouts formed a circle around the root-collar and lowest parts of the parent stems of the stool with the youngest sprouts at the periphery. The regeneration by seedlings was slight especially on the forest floor vegetation of the dwarf bamboo Sasa. The variety of size structure of stems and the existence of dead traces and/or dead center in each stool, the apparent difference in stool size, and the aggregations of stools in the forests suggest that stool expansion and long persistence of the stool at a given location may contribute to compensate for the scarcity of regeneration by seedlings inhibited by dwarf bamboo, and by the other shrubs and herbs.     

Diffused light conditions in canopy gaps in a beech (Fagus crenata Blume) forest

Summary A simple model for diffused light conditions in canopy gaps is presented. The vertical changes of illuminance in gaps calculated from the model agreed with the changes observed in a beech forest in Japan. The illuminance in a given sized gap in the forest could be estimated from the model. Size distribution of gaps in the forest suggested that most gaps were too small for the frequent occurrence of pioneer tree species.     

Structure of primary Japanese beech (Fagus japonica maxim.) forests in the Chichibu Mountains,central Japan,with special reference to regeneration processes

The floristic composition, structure and dynamics of three primaryFagus japonica stands were investigated in the Chichibu Mountains.F. japonica was dominant [RD(%): 64.9–87.0] and showed a slightly inverse J-shaped DBH class distribution in the quadrats [No. of canopy stems (H>20m): 87–138/ha]. The stems ofF. japonica for each size were distributed in the form of colonies, being scattered almost uniformly, and arranged in positive association with each other. Detailed examination of the bases of the stem groups forming colonies revealed that most of them originated from the bases of dead mother stems and that they were from common stools [No. of large stems (H>10 m) per stool: 6–11]. Among six major canopy gaps observed, only one included stems sprouting from the outer part ofF. japonica stools, while all the others were occupied by individuals of species other thanF. japonica. After tree-fall, several undercanopyF. japonica stems remained. Thus canopy gaps in these forest stands recovered through the sprouting of remainingF. japonica stools or by new sprouting ofF. japonica individuals adjacent to the gaps. However, it was considered difficult to fill canopy gaps only with sprouts when the distance between the center of a gap and that of a stool surpasses the crown vector. Such places that are not fully occupied by sprouts will be filled by individuals of other canopy and/or under-canopy species.     

Factors toxic to beech (Fagus sylvatica L.) seedlings in acid soils

The effects of highly and moderately acid soils on total biomass, biomass partitioning, fine root characteristics and nutritional status of beech seedlings (Fagus sylvatica L.) were studied in a growth chamber experiment. In Haplic Arenosols seedlings grew slowly but equally well without damage symptoms in a highly acid and a moderately acid soil horizon. The moderately acid A_h+B_w-horizon of a Eutric Cambisol was favourable to seedling growth. The fine root development was reduced in the highly acid A+B_w-horizon of a Dystric Cambisol and in the A_h+E-horizon of a Haplic Podzol, the latter of which also caused increased mortality. Seedling growth in the B₂-horizon of the Haplic Podzol was vigorous, in spite of a higher level of extractable Al and lower base saturation as compared with the A_h+E-horizon. These results are interpreted in relation to soil acidity, soil Al and nutritional status of the seedlings. We conclude that neither Al-toxicity nor nutrient deficiency cause the damage symptoms observed in the A_h+E-horizon of a Haplic Podzol and the fine root reduction in the A+B_w-horizon of a Dystric Cambisol. The damage symptoms of the PZh_A treatment seems to be more the result of H-toxicity or H-related factors other than nutrient shortage or Al-toxicity. Other pH-related toxic factors are discussed.     

A contribution to understanding the internal nitrogen budget of beech (Fagus sylvatica L.)

Summary The seasonal variation in the total nitrogen content of the xylem sap of the lower trunk section was investigated for two middle aged beech tree stands in northern Hessen each containing 130 trees. In addition seasonal changes in the percentage of nitrate in the total nitrogen content are described. The median values of the total nitrogen content of the xylem sap during the spring mobilization period reached 175 and 250 mg/l. During the summer about 35% of the total nitrogen in the xylem sap is in the form of nitrate. Finally, the distribution of NO₃ in the xylem sap along the trunk height was studied for two sample trees for each of the four seasons (n = 8).     

Forest development in canopy gaps in old-growth beech (Nothofagus) forests,New Zealand

Abstract. Tree size and age structures, treefall and canopy gap characteristics, and regeneration responses to treefalls were compared for three stands of old-growth beech (Nothofagus) forest dominated by N. fusca and N. menziesii on the South Island, New Zealand. Treefall gaps (up to 1000 m²) were most often caused by standing trees killed by drought and/or insect attack, or by trees snapped by wind. The causes of gap formation and the size and age distributions of treefall gaps varied between localities because of spatial and temporal differences in the histories of disturbance. At Fergies Bush where drought-related dieback had produced many large gaps with standing dead trees, gaps were generally young. Conversely, at Station Creek, small, old gaps formed by bolesnap dominated the disturbance regime. At Rough Creek, gaps of all ages and sizes were found along with an almost complete fern cover, and abundant shrubs and occasional subcanopy hardwood trees. Although overall patterns of regeneration were unrelated to differences in gap size, the relative abundance of N. fusca and N. menziesii varied between localities according to the seemingly minor differences in forest structure and disturbance history described above. Interpretations of regeneration response to gap parameters, therefore, need to account for differences in disturbance history between sites. Differences in the disturbance history between localities will also influence rates of gap closure, and because closure rates are used to estimate forest turnover times, meaningful comparisons of disturbance regimes for different forest types can only be made if this intersite variability is addressed.     

Effects of canopy gaps on the genetic structure of Camellia japonica saplings in a Japanese old-growth evergreen forest

The genetic structure of Camellia japonica saplings was investigated in relation to canopy conditions in an old-growth evergreen forest in Tsushima, Japan. To elucidate effects of canopy gaps on genetic structure, a 1 ha study site was divided into 20 x 20 m quadrats, which were classified into a gap quadrats (GAP), closed canopy quadrats (CLS) and mixed quadrats. Five GAP quadrats and six CLS quadrats were analyzed separately. Isolation-by-distance was tested by examining the correlation between genetic distance and geographic distance. A significant positive correlation was detected for GAP quadrats, whilst that for CLS quadrats was significantly smaller and not significantly different from zero. On the other hand, an analysis using Moran's I spatial autocorrelation coefficients indicates that the genetic structure is weaker in GAP quadrats than in CLS quadrats in short distance classes. The values were significantly positive for both types of quadrat. These results, along with our field observations on flowering, suggest that canopy gaps affect the genetic structure of C. japonica saplings in two distinct ways. First, canopy gaps may promote flowering and mating in an isolation-by-distance manner within canopy gaps. Second, canopy gaps may promote seed production and resulting overlap in seed shadows may weaken fine-scale genetic structures.     

High chloroplast haplotype diversity in Greek populations of beech (Fagus sylvatica L.)

The distribution of chloroplast DNA (cpDNA) variations in Greek beech ( Fagus sylvatica L.) populations was studied using chloroplast microsatellite markers. Thirteen haplotypes were identified from 40 populations by combining three different primers. Most of the cpDNA variation was distributed among populations, but a considerable variation was also observed within populations. The total diversity was very high for all regions. The N_st/G_st comparison was significant, indicating phylogenetic subdivision, but no strong spatial structure was detected, suggesting complex post-glacial migration patterns. Possible scenarios explaining this diversity pattern include the existence of several separated refugia in the region, the recolonisation of mountains by different beech lineages and the formation of an introgression zone between two different beech subspecies in the eastern part of the country.     

Differences in beech (Fagus crenata) regeneration between two types of Japanese beech forest and along a snow gradient

Differences in beech (Fagus crenata) regeneration were quantitatively investigated using power function analysis for the size–class (diameter at breast height, DBH) distribution and juvenile-to-canopy tree (J/C) ratio along a snow gradient throughout Japan. In snowy areas, all species combined, as well as F. crenata alone, showed constant regeneration, with parameter b≈−1.6 for the power function y=ax ^b (x=DBH, y=density), which is related to the DBH–class distribution. The good fit of the data to the function suggests that beech regenerates constantly with self-thinning patch dynamics. Parameter a, which indicates the abundance of small trunks, was high. Furthermore, the mean J/C ratio was ≈8, i.e., each parent beech tree produced eight juveniles. These results suggest that beech regenerates constantly with gap dynamics in snowy beech forests on the Japan Sea side of Japan (snowy). However, the fit of F. crenata was lower and nonsignificant in some forests in less snowy areas, despite the high fit of all species combined. In these areas, the mean of a was low, and b was often near zero for F. crenata regressions. These results suggest that the abundance of beech was low, and self-thinning was not evident because of the initial low abundance. Moreover, the mean J/C ratio was <1.0, suggesting that juvenile density was lower than that of canopy trees. Thus, the regeneration of F. crenata on the Pacific Ocean side of Japan (less snowy) is rather sporadic. Less snowy conditions may promote seed desiccation, predation of beechnuts and seedlings, and water stress. Lower F. crenata density may also reduce predator satiation and wind pollination.     

Potential risks for European beech (Fagus sylvatica L.) in a changing climate

Over large areas of Europe, coniferous monocultures are being transformed into mixed forests by the re-introduction of broadleaf tree species belonging to the potential natural vegetation. One important species of interest in this changing forest policy is European beech (Fagus sylvatica). However, at present, this forest management directive has ignored potential adverse effects of global climate change on wide-spread re-introduction of beech to these areas. Average global surface temperatures have risen by approx. 0.8°C in the period between 1861 and 2005 and are expected to continue to increase until the end of this century by 1.5–5.8°C above the 1990 value. To estimate the climate change in the southern part of central Europe in future, we reviewed calculations from regional climate models. Temperature increase for the southern part of central Europe is projected to be up to 2°C within the next 40 years. In contrast, the annual precipitation will most likely remain constant over the same time period, but will experience significant changes in seasonal patterns. Rising intensities of individual precipitation events may result in increasing number and intensities of flooding events and reduced precipitation during the growing season in a higher frequency of summer droughts. Growth and competitive ability of European beech will not, necessarily, respond to increasing CO₂ concentrations but may be strongly impacted by intensive drought that occurs during the growing season. Seedlings as well as adult trees may suffer from xylem embolism, restricted nutrient uptake capacity and reduced growth under limited water availability. However, it remains uncertain to what extent other environmental factors (e.g. soil properties, competitive interactions) may modify the drought response of beech, thus either enhancing susceptibility or increasing drought tolerance and resilience potential. Water-logged soils, predicted during the spring for several regions due to higher than average precipitation, could negatively impact nutrient uptake and growth of beech. Whereas other dominant species as, e.g. oak are well adapted to that environmental stress, beech is known to be sensitive to water-logging and flooding. Thus, the competitive capacity of beech might—depending on the other environmental conditions—be reduced under the expected future climate conditions. Silvicultural practices must be aware today of the potential risks which a changing climate may impose on sustainable forest development.     

Phylogeographical structure revealed by chloroplast DNA variation in Japanese beech (Fagus crenata Blume)

Intraspecific genetic variation in three non-coding chloroplast DNA (cpDNA) regions (trnT-L and trnL-F spacers, and trnL intron) of Japanese beech (Fagus crenata Blume) was investigated. This species is a major constituent of the typical cool-temperate deciduous forests in Japan. Twenty-one F. crenata populations from throughout Japan, and four F. japonica populations, a close relative of F. crenata, were examined. Seven haplotypes were distinguishable in F. crenata based on nucleotide substitutions and indels. Pairwise nucleotide diversities among haplotypes ranged from 0.0000 to 0.0042 for F. crenata, including F. japonica. The geographical distribution of cpDNA haplotypes was found to be highly structured in F. crenata. Four haplotypes predominated: haplotypes FC1 and FC4 are prevalent on the Pacific Ocean coast, haplotype FC6 is prevalent on the Japan sea coast from the San-in district to Hokkaido, whilst haplotype FC3 is restricted to northern Kyushu and the western-most part of Honshu. Two haplotypes (FC5 and FC7) are restricted to single populations and one haplotype (FC2) is a derivative of FC1. Each of these haplotypes, except FC2, are thought to be derived from different glacial refugia. Phylogenetic analysis showed that neither F. crenata nor F. japonica was monophyletic for the haplotypes, suggesting either ancestral polymorphism or ancient introgression between the lineages of these two Fagus species.     

Fluctuation of nut production and seedling appearance of a Japanese beech (Fagus crenata Blume)

The nut productivity, density of fallen nuts, seedling appearance and seedling survival of a Japanese beech (Fagus crenata Blume) were investigated at three localities, Mt Gozaisho, Mt Hodaka and Mt Bandai, Japan, from 1976 to 1992. Two patterns of cycles, a short cycle and a long one, were confirmed in the beech nut productivity. Synchronization in the long cycle was recognized both on Mt Hodaka and Mt Bandai. On Mt Gozaisho, the beech nut productivity was quite low, and the seedlings disappeared within 1 year. The phenomenon on Mt Gozaisho seemed to be caused by the low matter production mainly due to erosion and poor soils. A large number of seedlings appeared in the next spring of heavy mast years on Mt Hodaka and Mt Bandai. The large beech nut productivity contributed to the large seedling supply, and this enhanced the survival probability of beech seedlings. This demonstrates the possibility that beech seedlings survived longer even under dense dwarf bamboos, particularly if the seedling supply was large.     

Growth and nutrient dynamics of beech (Fagus sylvatica L.) seedlings in acid soils

The relative uptake rates of N, P, K, S, Ca, Mg, Fe, Mn, Zn, Cu, and Al were estimated in beech seedlings pot cultured in the field in six acid soils (treatments). The relative uptake rates were compared with the relative growth rates. The relative uptake rates of N, K and Ca agreed well with the growth rates of the seedlings irrespective of widely differing soil conditions (acid sand-clayey till, pH 4–6). The relative uptake rates of P, Fe, and Al differed from that predicted by the growth rate. The uptake rates of Fe and Al were highest at the lowest growth rates, and the P uptake rate was lower than the growth rate in these treatments. Thus the P availability probably limited growth in an eluvial (E) horizon of a podzol, and possibly in the illuvial (B) horizon of a podzol and in an acid clayey till (Dystric Cambisol). Low P uptake was associated with a tendency towards higher relative root growth rates. In terms of the concept of steady state nutrition the high relative root growth rate in some treatments may be interpreted as an acclimation to low P supply. The P limitation seemed to be related to interactions among Fe, Al and organic compounds of the soil solution.FAX no: +4646104423     

The effect of shoot architecture on hydraulic conductance in beech (Fagus sylvatica L.)

 Shoot hydraulic conductance was measured in beech (Fagus sylvatica L.) that had previously been exposed to high levels of nitrogen input. Whole-shoot hydraulic conductance, conductance per unit pressure gradient and leaf specific conductance were negatively correlated with the number of bud scars per unit length, a morphological parameter of tree decline. We propose a negative feedback mechanism by which stress induced alterations in shoot morphology can cause a lasting reduction of tree vigour. Received: 2 July 1997 / Accepted: 25 June 1998     

Developmental and seasonal changes of stress responsiveness in beech leaves (Fagus sylvatica L.)

The development of beech leaves (Fagus sylvatica L.) was characterized by determination of the pigment and electrolyte concentrations as well as the accumulation of dry mass and specific leaf mass from bud break to senescence. To test the hypothesis that stress tolerance and responsiveness of defences show developmental and/or seasonal changes, leaf discs were either incubated in the absence (control) or presence of paraquat to induce oxidative stress. Controls displayed developmental changes in stress susceptibility ranging from less than 15% of maximum electrolyte leakage in mature leaves to more than 20% leakage in senescent and 36–46% in immature leaves. Paraquat concentrations were chosen to result in about 95% of maximum electrolyte conductivity within 24 h in all developmental stages. Paraquat accumulation was about two‐fold lower in senescent as compared with immature leaves, whereas stress susceptibility, as characterized by the kinetics of the increase in relative leakage, was similar in these developmental stages with 50% of maximum electrolyte conductivity (EC₅₀) = 6·5 h in immature and 7·5 h in senescent leaves. In mature leaves with intermediate paraquat accumulation rates, two classes of stress‐sensitivity were distinguished, namely stress‐resistant and stress‐susceptible leaves with EC₅₀= 9·5 and 5·2 h, respectively. Stress‐resistance of mature leaves was accompanied by a rapid, approximately two‐fold induction of superoxide dismutase activity. Stress‐sensitive mature leaves initially contained high superoxide dismutase activities but showed a rapid, more than six‐ fold loss in activity in 24 h. Correlation of meteorological data with leakage rates suggested that high air temperatures and low precipitation might have been predisposing for loss of resistance against oxidative stress in beech leaves.     

Temperature dependency of bark photosynthesis in beech (Fagus sylvatica L.) and birch (Betula pendula Roth.) trees

Temperature dependencies of stem dark respiration (R(d)) and light-driven bark photosynthesis (A(max)) of two temperate tree species (Fagus sylvatica and Betula pendula) were investigated to estimate their probable influence on stem carbon balance. Stem R(d) was found to increase exponentially with increasing temperatures, whereas A(max) levelled off or decreased at the highest temperatures chosen (35-40 degrees C). Accordingly, a linear relationship between respiratory and assimilatory metabolism was only found at moderate temperatures (10-30 degrees C) and the relationship between stem R(d) and A(max) clearly departed from linearity at chilling (5 degrees C) and at high temperatures (35-40 degrees C). As a result, the proportional internal C-refixation rate also decreased non-linearly with increasing temperature. Temperature response of photosystem II (PSII) photochemistry was also assessed. Bark photochemical yield (Delta F/F(m)') followed the same temperature pattern as bark CO(2) assimilation. Maximum quantum yield of PSII (F(v)/F(m)) decreased drastically at freezing temperatures (-5 degrees C), while from 30 to 40 degrees C only a marginal decrease in F(v)/F(m) was found. In in situ measurements during winter months, bark photosynthesis was found to be strongly reduced. Low temperature stress induced an active down-regulation of PSII efficiency as well as damage to PSII due to photoinhibition. All in all, the benefit of bark photosynthesis was negatively affected by low (<5 degrees C) as well as high temperatures (>30 degrees C). As the carbon balance of tree stems is defined by the difference between photosynthetic carbon gain and respiratory carbon loss, this might have important implications for accurate modelling of stem carbon balance.     

Photoprotection mechanisms in European beech (Fagus sylvatica L.) seedlings from diverse climatic origins

This study shows that beech leaves adapt to their light environment by inducing dramatic changes to antioxidant systems and pigment composition. Thus, ascorbate, tocopherol, glutathione, β-carotene and xanthophyll cycle pigments are much more concentrated in sun leaves, while α-carotene is much less concentrated than in shade leaves. These characteristics were used to identify the inherent potential of beech cotyledons from three contrasting climatic origins to tolerate light stress. The antioxidant content was initially different in the three provenances tested, but these initial differences tended to reduce with leaf ageing. The higher antioxidant and de-epoxidized xanthophyll content found in developing cotyledons indicated a superior potential for tolerance to photo-oxidative damage in those plants collected from the stressful climate of the Pyrenees. Nevertheless under an experimental high irradiation treatment no differences in light stress tolerance were observed between provenances. Received: 31 May 1999 / Accepted: 16 November 1999